Biology and molecular phylogenetics of Nematoceras sulcatum, a second endemic orchid species from subantarctic Macquarie Island

A second orchid species, Nematoceras sulcatum M.A.Clem. et D.L.Jones, has been found on subantarctic Macquarie Island. A history of its discovery and recognition is provided. The morphology, biology and ecology of the new species are compared with N. dienemum (D.L.Jones) D.L.Jones, M.A.Clem. et Molloy, the other species of orchid on Macquarie Island. Molecular studies based on the internal transcribed spacer (ITS) region of nuclear ribosomal DNA reveal the phylogenetic relationship of the two Macquarie Island species compared to others in the genus from New Zealand and its Southern Ocean islands.     

The impact of introduced ship rats (Rattus rattus) on seedling recruitment and distribution of a subantarctic megaherb (Pleurophyllum hookeri)

Abstract The impact of introduced ship rats (Rattus rattus) on recruitment of the megaherb Pleurophyllum hookeri Buchan. (Asteraceae) was examined on subantarctic Macquarie Island, an island with no extant native terrestrial vertebrates. Pleurophyllum hookeri (Asteraceae) forms a dominant component of the Macquarie Island vegetation and is restricted to the subantarctic. The Macquarie Island population of P. hookeri is the most extensive and intact. Introduced ship rats (Rattus rattus) are well established in tall tussock grassland of Macquarie Island. We detected rat activity for the first time within P. hookeri herbfields, in autumn 2000. We found rats were destroying up to 90% of racemes. By excluding rats from caches of inflorescences that they had formed, we found they were having a significant negative effect on initial recruitment and seedling survival within the caches. However, because of high seedling mortality after 1 year, there was no sustained impact of the exclosures on P. hookeri seedling density.     

The first record of a terrestrial landhopper (Crustacea: Amphipoda: Talitridae) from Macquarie Island

A terrestrial talitrid crustacean is recorded for the first time from subantarctic Macquarie Island. The species, Puhuruhuru patersoni, is native to the southern part of the South Island of New Zealand. Its distribution on Macquarie Is. is restricted to a single locality near the ANARE station on the Isthmus. All mature females collected in December were carrying eggs; females appeared to be less mobile than males. It seems likely that the species has been transported accidentally from New Zealand, perhaps by the same means that introduced the isopod, Styloniscus otakensis, also from southern New Zealand.     

Warmer and less variable temperatures favour an accelerated plant phenology of two invasive weeds across sub‐Antarctic Macquarie Island

The great plasticity and diverse reproductive strategies of invasive alien plants are widely assumed to contribute to invasion success, even in extreme areas, often displacing native species. In this context, climate change creates new opportunities for biological invasions. Environmental variability and global warming are two of the climatic processes that may promote invasiveness, since alien species modulate their phenology to succeed under these circumstances. We monitored the phenological development (phenological stage advancement) of the two main invasive alien species: Poa annua L. and Cerastium fontanum Baumg. in the sub‐Antarctic Macquarie Island during the austral summer period along an altitudinal gradient. We found that higher temperatures lead to increased plant height and accelerated phenological development than lower temperatures in P. annua but found no direct evidence of the latter in C. fontanum. However, increased temperature variability negatively affected the phenological development of both species. Interestingly, despite their different reproductive strategy (rapid and impromptu in P. annua, and more synchronic and gradual in C. fontanum), both species prolifically succeeded in producing seeds at all sites showing the great acclimation of these two alien species even in limiting conditions. Since both alien species in Macquarie Island showed larger size and faster phenology at lower altitudes (i.e. milder conditions), this would indicate a great influence of ameliorating abiotic extremes on alien plant invasive capabilities at environmental extremes. Thus, our results warn of the increasing capabilities under climatic warming scenarios for alien plants to reproduce even at such remote ranges. This highlights the need to reinforce calls for special attention to prevent the spread of these kinds of species to other similar sub‐polar areas, where intensive post‐introduction management may be difficult or expensive.     

Long distance migration of insects to a subantarctic island

Transoceanic migration of four species of macrolepidoptera to subantarctic Macquarie Island has been detected in 7 out of 33 years during the period 1962–96 and is restricted to spring and autumn. Analyses of synoptic charts during the migration period show that autumn immigrants originated from New Zealand and comprised a single species of noctuid moth,Agrotis ipsilon (Walker). Spring immigrants originated from Australia and comprised two noctuids, Dasypodia selenophora Guenée and Persectania ewingii Westwood and a butterfly, Vanessa kershawi (McCoy). Autumn migrations were associated with depressions in the southern Tasman Sea. Spring migrations were associated with the eastward passage of prefrontal airflows ahead of cold fronts which extended from southern Australia to the west of Macquarie Island. In an analysis of one of these events, winds exceeded 30 ms^?1 at 300 m altitude and could have transported migrants from Tasmania to Macquarie Island overnight in less than 10 h. Flight activity was assisted by the presence of a nocturnal temperature inversion that maintained upper air temperatures above 5 °C. The effect of potential global warming on the migration and colonization of Macquarie Island by insects is discussed.     

Phenotypic plasticity and genetic diversity in Poa annua L. (Poaceae) at Crozet and Kerguelen Islands (subantarctic)

The widely distributed grass, Poa annua, is one of the most common alien species in the subantarctic islands. The historical events of its introduction remain generally unknown, as well as the evolutionary consequences of its colonisation in these remote environments. Populations from the Crozet archipelago and Kerguelen Islands were compared in terms of morphology, cytogenetics and enzyme polymorphism. Seeds from natural populations were also sown in an experimental garden in France to test phenotypic plasticity. This preliminary study demonstrated the high phenotypic plasticity in P. annua in the French subantarctic islands. This plasticity and allotetraploidy could be important factors which reinforce the colonising capacities of P. annua. Our results revealed the low genetic diversity of the populations analysed, which could be related to the founding effect or to the fragmentation of the populations. Accepted: 25 April 1999     

Fur seals at Macquarie Island: post-sealing colonisation,trends in abundance and hybridisation of three species

Commercial sealers exterminated the original fur seal population at Macquarie Island in the early 1800s. The first breeding record since the sealing era was not reported until March 1955. Three species of fur seal now occur at Macquarie Island, the Antarctic (Arctocephalus gazella), subantarctic (A. tropicalis) and New Zealand (A. forsteri) fur seal. Census data from 54 breeding seasons in the period 1954–2007 were used to estimate population status and growth for each species. Between the 1950s and 1970s, annual increases in pup production for the species aggregate were low. Between 1986 and 2007, pup production of Antarctic fur seals increased by about 8.8% per year and subantarctic fur seals by 6.8% per year. The New Zealand fur seal, although the most numerous fur seal species on Macquarie Island, has yet to establish a breeding population, due to the absence of reproductively mature females. Hybridisation among species is significant, but appears to be declining. The slow establishment and growth of fur seal populations on Macquarie Island appears to have been affected by its distance from major population centres and hence low immigration rates, asynchronous colonisation times of males and females of each species, and extensive hybridisation.     

Lepidoderma cristatosporum, a new species of myxomycete from Australia

A new species of myxomycete, Lepidoderma cristatosporum, is described from Australia. This species is represented by material collected on subantarctic Macquarie Island. Photographs of fruiting bodies and morphological features as observed under light microscopy are provided, along with micrographs of spore ornamentation as observed by scanning electron microscopy. In addition, the new species is compared with other taxa belonging to the genera Lepidoderma and Diderma, especially with the type material of Lepidoderma crustaceum, a very similar species, and Diderma ochraceum and D. sauteri, two species also muscicolous, for which microphotographs are provided.     

A new species of Didymium (Myxomycetes) from subantarctic Macquarie Island

A series of specimens of a stalked Didymium collected during an intensive survey of fungal biodiversity on subantarctic Macquarie Island were determined to represent a species new to science. The new species Didymium macquariense is described and compared with other morphologically similar species. Micrographs of the most important morphological characters are provided.     

Barging in: A Temperate Marine Community Travels to the Subantarctic

A diverse fouling community discovered encrusting a barge intended for deployment at subantarctic Macquarie Island is described and its role as a transport vector for non-indigenous marine organisms is discussed. The barge proved to be a potential vector capable of transporting entire epi-bethic communities, 20 species in total, from a temperate estuarine system (Derwent River, Tasmania, Australia) into the subantarctic. For one invasive amphipod species Monocorophium acherusicum, over 136000 individuals including ovigerous females were calculated to be associated with the barge fouling community. Although distinct differences exist between the thermal ranges of Macquarie Island and the Bruny bioregion of Tasmania, a hazard assessment based on the Gower similarity coefficient suggested sufficient similarity between the two environments to allow for survival of transported organisms for eight months of the year. Several invasive species are able to survive the thermal conditions of the subantarctic irrespective of the time of year. This study identifies the need for effective quarantine measures aimed at identifying and managing marine biosecurity hazards in association with human activities in high latitude regions.     

Hidden diversity in high-latitude Southern Hemisphere environments: Reinstatement of the genus Rama and description of Vandenhoekia gen. nov. (Cladophoraceae,Ulvophyceae, Chlorophyta), two highly variable genera

The continental coasts and remote islands in the high-latitude Southern Hemisphere, including the subantarctic region, are characterized by many endemic species, high abundance of taxa, and intermediate levels of biodiversity. The macroalgal flora of these locations has received relatively little attention. Filamentous green algae are prolific in the intertidal of southern islands, but the taxonomy, distribution, and evolutionary history of these taxa are yet to be fully explored, mostly due to the difficulty of access to some of these locations. In this study, we examined specimens of the order Cladophorales from various locations in the high-latitude Southern Hemisphere including the subantarctic (the Auckland Islands, Bounty Islands, Campbell Island, Macquarie Island, and Kerguelen Islands), as well as mainland New Zealand, the Chatham Islands, Chile, and Tasmania. The analyses of the rDNA sequences of the samples revealed the existence of two new clades in a phylogeny of the Cladophoraceae. One of these clades is described as the novel genus Vandenhoekia gen. nov., which contains three species that are branched or unbranched. The amended genus Rama is reinstated to accommodate the other clade, and contains four species, including the Northern Hemisphere “Cladophora rupestris.” In Rama both branched and unbranched morphologies are found. It is remarkable that gross morphology is not a predictor for generic affiliations in these algae. This study illustrates that much can still be learned about diversity in the Cladophorales and highlights the importance of new collections, especially in novel locations.     

Subantarctic flowering plants: pre‐glacial survivors or post‐glacial immigrants?

Aim The aim here was to assess whether the present‐day assemblage of subantarctic flowering plants is the result of a rapid post‐Last Glacial Maximum (LGM) colonization or whether subantarctic flowering plants survived on the islands in glacial refugia throughout the LGM. Location The circumpolar subantarctic region, comprising six remote islands and island groups between latitudes 46° and 55° S, including South Georgia in the South Atlantic Ocean, the Prince Edward Islands, Îles Crozet, Îles Kerguelen, the Heard Island group in the South Indian Ocean and Macquarie Island in the South Pacific Ocean. Methods Floristic affinities between the subantarctic islands were assessed by cluster analysis applied to an up‐to‐date dataset of the phanerogamic flora in order to test for the existence of provincialism within the subantarctic. A review of the primary literature on the palaeobotany, geology and glacial history of the subantarctic islands was carried out and supplemented with additional palaeobotanical data and new field observations from South Georgia, Île de la Possession (Îles Crozet) and Îles Kerguelen. Results First, a strong regionalism was observed, with different floras characterizing the islands in each of the ocean basins, and endemic species being present in the South Indian Ocean and South Pacific Ocean provinces. Second, the majority of the plant species were present at the onset of accumulation of post‐glacial organic sediment and there is no evidence for the natural arrival of new immigrants during the subsequent period. Third, a review of geomorphological data suggested that the ice cover was incomplete during the LGM on the majority of the islands, and ice‐free biological refugia were probably present even on the most glaciated islands. Main conclusions Several independent lines of evidence favour the survival of a native subantarctic phanerogamic flora in local refugia during the LGM rather than a post‐LGM colonization from more distant temperate landmasses in the Southern Hemisphere.     

Effects of temperature on growth rates of fungi from subantarctic Macquarie Island and Casey,Antarctica

Summary The linear growth rates of fungal isolates were measured on agar plates at temperatures ranging from 4° to 35°C. Fungi tested included the major fungal colonizers of leaves and litter of the three dominant plant species on subantarctic Macquarie Island, and major fungal species associated with plant and soil communities near Australia's Casey Station on the Antarctic Continent. All fungi grew at 4°C and were classified as psychrotrophs. Maximum growth rates were recorded at temperatures of 10° to 20°C for 13 of the 15 isolates from Macquarie Island and for all six isolates from Casey. Most of the leaf colonizing fungi from Macquarie Island had optimum growth temperatures of 15°C whereas all litter fungi from Macquarie Island and Casey fungi except Thelebolus microsporus had optimum growth temperatures of 20°C or above. Maximum growth of all species was at temperatures above those normally prevailing in their natural environments, with most species growing at 4°C at between 10% and 30% of their maximum rates. However, microclimatic effects may have resulted at times in temperatures near their growth optima. The highest growth rates at 4°C were recorded for Phoma spp. 1 and 2, Phoma exigua and Mortierella gamsii from Macquarie Island and Mortierella sp. 1 from Casey. Thelebolus microsporus and sterile sp. G from Casey also grew relatively fast at 4°C, and these species, and Phoma sp. 3 and Phoma exigua from Macquarie Island had the lowest Q-₁₀ values for the temperature range 4° to 15°C.     

Changes in the cover of plant species associated with climate change and grazing pressure on the Macquarie Island coastal slopes, 1980–2009

Climate change and alien species have affected the vegetation of subantarctic islands. Long-term monitoring of vegetation change on the steep coastal slopes of subantarctic Macquarie Island has allowed responses of plant species to various disturbance regimes to be well documented, although, until recently, the confounding effect of feral herbivore disturbance obscured any responses that might be attributed to climate change. The uncoupling of climate change from variation in feral rabbit numbers allowed us to test whether any plant species were increasing or decreasing on the coastal slopes of the island between 1980 and 2009, independent of rabbit grazing pressure. We used analysis of variance to test for differences in species cover classes between four measurement times on each of 101 quadrats in each of 1980/1981, 1995, 2003 and 2009. We had 54 quadrats on landslips and 47 elsewhere. Approximately two-thirds of the species with significant temporal change exhibited changes that could be expected from variation in rabbit grazing pressure. However, approximately one-third of the species increased in cover irrespective of grazing pressure. On landslips, variation in the cover of these increaser species was largely related to time in a linear mixed model, whereas elsewhere altitude and time were both important. The increase in both atmospheric dryness and episodic soil water-logging that has been described for the island since 1980 may best explain the increaser species.     

DISTINCTION OF SOUTH AMERICAN HAPLOGLOIA ANDERSONII AND CHORDARIA LINEARIS (PHAEOPHYCEAE) BASED ON DIFFERENCES IN SEXUAL REPRODUCTION,MORPHOLOGY, ECOPHYSIOLOGY,AND DISTRIBUTION1

Haplogloia andersonii (Farlow) Levring is an anti-tropical species that occurs on cold and warm-temperate Pacific coasts of both Americas. In its habit it resembles the subantarctic species Chordaria linearis (Hooker et Harvey) Cotton. Culture studies show that the species differ in morphology and ecophysiology of their microscopic gametophytes and in gamete behavior. Details of sporophyte anatomy are presented that also allow the distinction of field plants. In South America, H. andersonii occurs only on the Pacific coast, from central Perú (14°S) to southern Chile (50°S). Chordaria linearis occurs on the Pacific coast from Chiloé Island (43°S) to Cape Horn (56°S). In the shared area the species may co-occur. On the Atlantic coast, C. linearis was newly collected at a locality in northern Patagonia (41°S). In addition, C. linearis occurs in Antarctica. Haplogloia moniliformis Richer, recently described from Macquarie Island, is probably synonymous with Chordaria linearis.     

The invertebrates of sub-Antarctic Bishop Island

We report on the first comprehensive collection of invertebrates made on sub-Antarctic Bishop Island, which lies 33 km south of Macquarie Island. A total of 15 species were collected, of which 14 also occur amongst the 168 species recorded from Macquarie Island. We hypothesise that the greater species richness of Macquarie Island relative to Bishop Island is largely caused by the greater habitat diversity on Macquarie Island but that other factors, including accidentally introduced species, may also be important. Received: 3 July 1996 / Accepted: 2 October 1996     

Curvilinear Effects of Invasive Plants on Plant Diversity: Plant Community Invaded by Sphagneticola trilobata

The effects of invasive plants on the species diversity of plant communities are controversial, showing either a positive or negative linear relationship. Based on community data collected from forty 5 m×5 m plots invaded by Sphagneticola trilobata in eight cities across Hainan Island, China, we found S. trilobata decreased plant community diversity once its cover was beyond 10%. We demonstrated that the effects of invasive/native plants on the plant diversity of communities invaded by S. trilobata were curvilinear. These effects, which showed peaks under different degrees of vegetation cover, appeared not only for S. trilobata and all invasive plants, but also for all native plants. Invasive plants primarily had negative effects on plant diversity when they became abundant at a much lower cover level (less than 35%), compared with the native plants (over 60%). Thus, it is necessary to distinguish a range for assessing the effects of plants, especially invasive plants. Our results also confirmed that the invasion intensity of invasive alien plants increased with the intensity of local economic development. We highlight and further discuss the critical importance of curvilinear effects of biological invasion to provide ideas regarding the conservation of local biodiversity and the management of invasive plants.     

High levels of genetic variability in the moss Ceratodon purpureus from continental Antarctica, subantarctic Heard and Macquarie Islands, and Australasia

The random amplified polymorphic DNA (RAPD) technique, and DNA sequencing of the conserved nuclear ribosomal DNA internal transcribed spacer region (ITS1-5.8S-ITS2), have been used to assess levels of genetic diversity in the moss Ceratodon purpureus from several locations in Australasia, subantarctic Heard and Macquarie Islands, and continental Antarctica. Populations from Heard and Macquarie Islands and from Antarctica maintain high levels of genetic variation. Both within- and among-colony variation were observed at these locations. DNA sequence analysis showed that samples from the Ross Sea region of Antarctica were most closely related to colonies from Casey and Macquarie Island, and that one colony from Heard Island was most closely related to one from Europe. DNA sequence data separated two Australian populations from the Antarctic and subantarctic group on a dendrogram. Detailed RAPD analysis of a single colony from continental Antarctica demonstrated that mutation probably causes the high variability observed in this moss. DNA sequencing and RAPD analysis are complementary techniques for genetic investigation of Antarctic moss populations.Jenny Ninham was an integral part of this research team for several years. Unfortunately she did not live to see these results published.     

The influence of landscape,climate and history on spatial genetic patterns in keystone plants (Azorella) on sub‐Antarctic islands

The distribution of genetic variation in species is governed by factors that act differently across spatial scales. To tease apart the contribution of different processes, especially at intermediate spatial scales, it is useful to study simple ecosystems such as those on sub‐Antarctic oceanic islands. In this study, we characterize spatial genetic patterns of two keystone plant species, Azorella selago on sub‐Antarctic Marion Island and Azorella macquariensis on sub‐Antarctic Macquarie Island. Although both islands experience a similar climate and have a similar vegetation structure, they differ significantly in topography and geological history. We genotyped six microsatellites for 1,149 individuals from 123 sites across Marion Island and 372 individuals from 42 sites across Macquarie Island. We tested for spatial patterns in genetic diversity, including correlation with elevation and vegetation type, and clines in different directional bearings. We also examined genetic differentiation within islands, isolation‐by‐distance with and without accounting for direction, and signals of demographic change. Marion Island was found to have a distinct northwest–southeast divide, with lower genetic diversity and more sites with a signal of population expansion in the northwest. We attribute this to asymmetric seed dispersal by the dominant northwesterly winds, and to population persistence in a southwestern refugium during the Last Glacial Maximum. No apparent spatial pattern, but greater genetic diversity and differentiation between sites, was found on Macquarie Island, which may be due to the narrow length of the island in the direction of the dominant winds and longer population persistence permitted by the lack of extensive glaciation on the island. Together, our results clearly illustrate the implications of island shape and geography, and the importance of direction‐dependent drivers, in shaping spatial genetic structure.     

Native species richness buffers invader impact in undisturbed but not disturbed grassland assemblages

Many systems are prone to both exotic plant invasion and frequent natural disturbances. Native species richness can buffer the effects of invasion or disturbance when imposed in isolation, but it is largely unknown whether richness provides substantial resistance against invader impact in the face of disturbance. We experimentally examined how disturbance (drought/burning) influenced the impact of three exotic invaders (Centaurea stoebe, Linaria dalmatica, or Potentilla recta) on native abundance across a gradient of species richness, using previously constructed grassland assemblages. We found that invaders had higher cover in experimentally disturbed plots than in undisturbed plots across all levels of native species richness. Although exotic species varied in cover, all three invaders had significant impacts on native cover in disturbed plots. Regardless of disturbance, however, invader cover diminished with increasing richness. Invader impacts on native cover also diminished at higher richness levels, but only in undisturbed plots. In disturbed plots, invaders strongly impacted native cover across all richness levels, as disturbance favoured invaders over native species. By examining these ecological processes concurrently, we found that disturbance exacerbated invader impacts on native abundance. Although diversity provided a buffering effect against invader impact without disturbance, the combination of invasion and disturbance markedly depressed native abundance, even in high richness assemblages.